A RECENT SURVEY OF THE AMPHIBIAN FAUNA OF THE CARDAMOM...

ITHE RAFFLES BULLETIN OF ZOOLOGY 2002 50(2): 465-481@ National University of Singapore

A RECENT SURVEY OF THE AMPHIBIAN FAUNA OF THE CARDAMOM MOUNTAINS,SOUTHWEST CAMBODIA WITH DESCRIPTIONS OF THREE NEW SPECIES

Annemarie OhlerLaboratoire des Reptiles et Amphibiens, Museum National d'Histoire Naturelle, 25 rue Cuvier, F-75005 Paris, France

Steven R. SwanFauna and Flora International, Indochina Programme Office, PO Box 78, I04B Pho Hue, Hanoi, Vietnam

Jennifer C. DaltryFauna and Flora International, Great Eastern House, Tenison Road, Cambridge CBI 21T, UK

ABSTRACT. -Reported here are the findings of the first field surveys for amphibians in the CardamomMountains of Southwest Cambodia, which were conducted in January -March 2000 and February -April2001, A total of34 species of anuran were found in these mountains, including new species of Megophrys,Philautus and Rana (Sylvirana) that are described here. A further 17 species that have not previously beenreported from Cambodia were also recorded during these surveys. These results double the size ofCambodia's known amphibian fauna. Species of particular taxonomic interest from the collections arediscussed in further detail, and a national checklist of Cambodia's known amphibian fauna is also provided.

KEY WORDS. -Amphibians, new species, Megophrys, Philautus, Rana (Sylvirana), Cambodia, taxonomy.

iI

INTRODUCTION The Cardamom Mountains are the dominant topographicalfeature of Southwest Cambodia, occupying an area of over

The existing knowledge of the amphibian fauna of Indochina, one million hectares (10,000 km2). The mountain range hasin terms of both composition and distribution, is highly an axis orientated Northwest-Southeast running forfragmented. The incompleteness of our knowledge is approximately 200 km, and an overall altitudinal variationreflected in the continual discovery of new species from of c. 100 to 1,771 m above sea level- Phnom [Mount] Aural,within the region. The past decade has witnessed an Cam?odia's highest mountain. The mountains inclu~e t,:"ointensification of research interest in the Indochinese nomInally protected areas: the Phnom Samkos WIldlifepeninsula, yet only in the last two years have the scattered Sanctuary and the Phnom Aural Wildlife Sanctuary.

data on amphibian distribution and taxonomy been collated Th C d M . h . d b h. h . f"11e ar amom ountains are c aractenze y ig rain illto provide some nation-wide reviews, e.g., Stuart (1999) for ., h f th .th .. th.lor muc 0 e year, Wi some areas reCeiVIng more anLaos; Inger et al. (1999) for Vietnam. However, of all the 4 000 f .11 (A h 11 1997) aki th.., mm 0 rain annua y s we , , m ng eIndochinese nations, Cambodia has received the least C d M t . th .. t 1 . C b d .

..ar amom oun ams e ramies pace m am 0 iaattention from batrachologists. (Weiler, 1998). Humidity and temperature are also high with

slight seasonal variation (Ashwell, 1997). TemperaturesNot much is known of Cambodian amphibians because range from between 25°C and 30°C on average, although atduring French occupation little attention was paid to higher elevations temperatures can drop below 15°C at nightdocumenting the colony's fauna and a recent history of war (Momberg & Daltry, 2000). The dry season lasts fromhas hindered any modern scientific investigation in the field. December to March and the rainy season from May toThis conflict persisted up until the very end of the last century October when the mountains capture the Southwest monsoonand only in the past three years has security stabilised and arriving offshore from the Gulf of Thailand.improved sufficiently to allow relatively safe exploration ofCambodia's forests. Until the present survey, only 18 species Three distinct basic vegetation types, that are principally awere confirmed as present in Cambodia (see Bourret, 1942, function of altitude, cover the Cardamom Mountains. Dryfor a review; van Dijk, unpublished data), most of these being dipterocarp forest occupies the lowland basins between thecommon and widely distributed throughout Southeast Asia. mountains, hill evergreen forest covers the lower slopes of

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Ohler et al.: Amphibia of Cambodia

Table I. Sites surveyed for amphibians in the Cardamom Mountains 2000-2001.

Site Location Province Geo-coordinates Duration

Phnom Samkos Wildlife Western Cardamom Pursat 120 15'N 1030 OO'E 4 weeksSanctuary Mountains (24/1-27/2/00)

T'Mar Bang District Central Cardamom Koh Kong 110 50'N 1030 35'E 4 weeksMountains (25/2-27/3/00)

Phnom Aural Wildlife Eastern Cardamom Kampong Speu 120 OO'N 1040 10'E 5 weeksSanctuary Mountains (10/2-21/3/01)

Veal Veng wetland Central Cardamom Pursat 120 05'N 1030 15'E I weekMountains (30/3-6/4/01)

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the mountains up to 1,200 m, and montane evergreen forest at night, usually between 19:00 and 23:00 hours. Because

continues above 1,200 m on the upper slopes and mountain this work was carried out as part of a wider conservation

summits. Together, the hill and montane evergreen forests initiative, only a limited number of voucher specimens were

of Southwest Cambodia form one of the distinct bio-climatic taken (1 to 10 specimens) to represent the full range of

regions recognised for the country -the Cardamom species encountered. Frogs were caught by hand and

evergreen forest eco-region (Fontanel, 1972). specimens fixed in c. 4% formalin and stored in c. 70%

ethanol. Voucher specimens from our surveys are depositedThe present paper summarises the findings of amphibian in BMNH (Natural History Museum, London, United

surveys of the Cardamom Mountains in the Southwest of Kingdom) and MNHN (Museum, national d'Histoire

Cambodia. The surveys were conducted during the dry naturelle, Paris, France) (see Tab1e2). Historical museum

seasons of 2000 (January to March) and 2001 (February to material examined for this study is listed as an Appendix.

April), as part of a larger biodiversity study of these remote Specimens were measured using callipers and a graded rule

mountains requested by the Cambodian Government for the under a binocular microscope. In order to facilitate

purposes of conservation management planning (Daltry & comparisons, the description's methodology and plan were

Momberg, 2000). As the most extensive amphibian surveys the same as those used in previous works on Asian anuransto have been carried out in Cambodia to date, our work makes (Dubois & Ohler, 1998, 1999, 2000; Ohler & Dubois, 1999;

a significant contribution to the existing knowledge of Ohler et al., 2000; Veith et al., 2001). The webbing formula

amphibian distributions in Indochina. is given according to Myers & Duellman (1982). The

relevant synonymy for each species reported on is listed in

Table 8.

METHODSThe present paper cites several recent records based on the

Amphibians were surveyed in four localities within the unpublished data of Peter Paul van Dijk, who is currently

Cardamom Mountain range, details of which are given in preparing a field guide to the amphibian species of Southeast

Table 1. During the 2000 dry season, the Phnom Samkos Asia. Through van Dijk's kind generosity, we were able to

Wildlife Sanctuary and T'Mar Bang District, Koh Kong include the specimens he has studied in Table 8. Other data

Province, in the Central Cardamom Mountains were come from various authors in particular from Bourret. In

surveyed. The Phnom Samkos Wildlife Sanctuary occupies 1942 he published an important book on the frog fauna of

3,338 km2 of the western end of the Cardamom Mountain southeast Asia, the only consistent review even today.

range. Phnom Samkos itself is the highest point in the However, it included little new information, as most of the

sanctuary and the second highest in the country attaining an data (including Bourret's own data) had already been

altitude of 1,717 m. The highest elevation of the T'Mar Bang published elsewhere (see Bourret, 1942). Therefore, while

survey area in the Central Cardamom Mountains is c. 1,200 Bourret (1942) is an excellent source, it nevertheless remains

m and the area is located within a newly proposed protected a secondary source, and we prefer instead to cite primary

area. However, at the time of the survey, the second site sources for the relevant synonymies (see Table 8) and cite

was under logging concession. During 2001, the Phnom this book only when original data are published there.

Aural Wildlife Sanctuary was the focus of field surveys. The

Phnom Aural Wildlife Sanctuary encompasses 2,537 km2 Voucher specimens were deposited in BMNH (Natural

of the far eastern end of the Cardamom Range including History Museum, London, United Kingdom) and MNHN

Cambodia's highest mountain, Phnom Aural at 1,771m. One (Museum National d'Histoire Naturelle, Paris, France).

week was spent at the fourth study site, the Veal Veng

wetland, which lies between the Phnom Samkos Wildlife The following abbreviations are used for measurements:

Sanctuary and the second survey site in T'Mar Bang District. SVL: Snout-vent length. Head: HW: Head width; HL: Head

The wetland is a highly modified mosaic of inundated length (from the back of the mandible to the tip of snout);

habitats at an elevation of 560 m. MN: Distance from the back of the mandible to the nostril;

MFE: Distance from the back of the mandible to the frontThe majority of survey effort was focused on aquatic habitats of the eye; MBE: Distance from the back of the mandible

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THE RAFFLES BULLETIN OF ZOOLOGY 2002

Table 2. List of amphibian species encountered during the 2000-200 I surveys of the Cardamom Mountains, Pursat, Koh Kong and KampongSpeu Provinces, Southwest Cambodia.

SPECIES Number of Collection numberspecimens

MEGOPHRYIDAE Bonaparte, 1850

Leptolalax sp. I MNHN 2001.0204Megophrys (Xenophrys) auralensis, new species 10 BMNH 2000.0077, MNHN 2001.0205-0213

BUFONIDAE Gray, 1825 !

Bufo macrotis Boulenger, 1887 3 BMNH 2000.0074-0075, MNHN 2001.0201Bufo melanostictus Schneider, 1799 2 MNHN 2001.0202-0203Bufo parvus Boulenger, 1887 2 BMNH 2000.0072-0073

RANIDAE Rafinesque-Schmaltz, 1814

Chirixalus doriae Boulenger, 1893 4 MNHN 2001.0219-0222Chirixalus vittatus (Boulenger, 1887) 2 BMNH 2000.0144, MNHN 2000.0223Fejervarya limnocharis (Gravenhorst, 1829) 4 BMNH 2000.0108-0111Hoplobatrachus chinensis (Osbeck, 1765) 2 BMNH 2000.0092-0093Limnonectes (Elachyglossa) gyldenstolpei 7 BMNH 2000.0112-0114, BMNH 2000.0125, MNHN(Anderson, 1916) 2001.0224-0226Limnonectes (Elachyglossa) kohchangae 18 BMNH 2000.0115-0124, MNHN 2001.0227-0234(Smith, 1922)Occidozyga lima (Gravenhorst, 1829) 4 BMNH 2000.0094-0096, MNHN 2001.0235Paa (Eripaa)fasciculispina (Inger, 1970) 8 BMNH 2000.0102-0107, MNHN 2001.0236-0237Philautus cardamonus, new species 3 BMNH 200.0149-0151Philautus parvulus (Boulenger, 1893) 7 BMNH 2000.0145-0148, MNHN 2001.0238-0240Phrynoglossus martensii Peters, 1867 8 BMNH 2000.0097-0101, MNHN 2001.0241-0243Polypedates cf. leucomystax (Gravenhorst, 1829) 3 BMNH 2000.0152-0154Rana (Hylarana) erythraea (Schlegel, 1837) 4 BMNH 2000.0126, MNHN 2001.0244-0246Rana (Hylarana) macrodactyla (Gunther, 1859). 5 BMNH 2000.0127-0128, MNHN 2001.0247-0249Rana (Hylarana) taipehensis van Denburgh, 1909 I BMNH 2000.0129Rana (Sylvirana) faber, new species 15 BMNH 2000.0134, BMNH 2000.0136-0141, MNHN

2001.0256-0263Rana (Sylvirana) mortenseni Boulenger, 1903 9 BMNH 2000.0133, BMNH 2000.0135, BMNH 2000.0143,

MNHN 2001.0250-0255Rhacophorus bipunctatus Ahl, 1927 3 BMNH 2000.0157-0159Rhacophorus bisacculus Taylor, 1962 2 BMNH 2000.0155-0156Theloderma asperum (Boulenger, 1886) I MNHN 2001.0264

MICROHYLIDAE Gunther, 1858

Kalophrynus interlineatus Blyth, 1855 I MNHN 2001.0214Kaloula pulchra Gray, 1831 I BMNH 2000.0078Micryletta inornata (Boulenger, 1890) I BMNH 2000.0079Microhyla annamensis Smith, 1923 1 BMNH 2000.0086Microhyla berdmorei (Blyth, 1856) 3 BMNH 2000.0080-0082Microhyla butleri Boulenger, 1900 7 BMNH 2000.0089-0091, MNHN 2001.0215-0218 :Microhyla heymonsi Vogt, 1911 1 BMNH 2000.0083Microhyla ornata (Dumeril & Bibron, 1841) 2 BMNH 2000.0087-0088Microhyla pulchra (Hallowell, 1861) 2 BMNH 2000.0084-0085

to the back of the eye; lFE: Distance between the front of fingers I to IV; W AI -IV Width of fingers I to IV. Hindlimb:the eyes; IBE: Distance between the back of the eyes; IN: FL: Femur length (from vent to knee); TL: Tibia length; FOL:Internasal space; EN: Distance from the front of the eye to Foot length (from the base of the inner metatarsal tuberclethe nostril; EL: Eye length; SN Distance from the nostril to to the tip of the toe); FTL: Fourth toe length (from the basethe tip of the snout; SL Distance from the front of the eye of the first subarticular tubercle); PPI -V Width of pads ofto the tip of the snout; TYD: Greatest tympanum diameter; toes I to V; WPI -V Width of toes I to V; IMT: Length ofTYE: Distance from tympanum to the back of the eye; IUE: inner metatarsal tubercle; ITL: Inner toe length. Webbing:Minimum distance between upper eyelids; UEW: Maximum MTTF: Distance from the distal edge of the metatarsalwidth of inter upper eyelid. Forearm: HAL: Hand length tubercle to the maximum incurvation of the web between(from the base of the outer palmar tubercle to the tip of the third and fourth toe; TFTF: Distance from the maximumtoe); FLL: Forelimb length (from the elbow to the base of incurvation of the web between third and fourth toe to thethe outer tubercle); TFL: Third finger length (from the base tip of fourth toe; MTFF: Distance from the distal edge ofof the first subarticular tubercle); PAl -IV Width of pads of the metatarsal tubercle to the maximum incurvation of the

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web between fourth and fifth toe; FFTF: Distance from the diameter of eye (EL 9.21 mm). (4) Canthus ro~tralis rathermaximum incurvation of the web between fourth and fifth sharp, loreal region concave, acute. (5) Interorbital spacetoe to the tip of fourth toe; WTF: Webbing between third slightly convex, larger (IUE 8.55 mm) than upper eyelidand fourth toe (from the base of the first subarticular (UEW 6.84 mm) and internarial distance (IN 8.16 mm);tubercle); WFF: Webbing between fourth and fifth toe (from distance between front of eyes (IFE 11.3 mm) about twothe base of the first subarticular tubercle); WI: Webbing times the distance between back of eyes (IBE 20.7 mm).between third and fourth toe when folded along fourth toe (6) Nostrils oval with low flap of skin laterally, closer to(from the base of the first subarticular tubercle); WII: eye (EN 2.76 mm) than to tip of snout (NS 4.47 mm). (7)Webbing between fourth and fifth toe when folded along Pupil indistinct. (8) Tympanum (TYD 6.05 mm) distinct,fourth toe (from the base of the first subarticular tubercle). oval, oblique, 60% of eye diameter; tympanum-eye distance

(TYE 6.71 mm) 110% of tympanum diameter. (9) PinealOther abbreviations -p. t.: per thousand. ocellus absent. (10) Vomerine ridge present, short, rounded,

without teeth, parallel to body axis, closer to choanae thanfrom each other, shorter than distance between them. (11)

SYSTEMATICS Tongue large, oval, entire, bearing no median lingual process.Tooth-like projections on lower jaw absent. (12)

A total of 34 species were recorded during the four months Supratympanic fold ?istinct, from 'back of eye to a.bovespent in the field. These species are listed in Table 2. Three shoulder. (13) Parot~Id glan?s absent. (14) Cephalic ndgesf th ' t . d 17 th .absent. (15) Co-ossIfied skin on head absent.0 e speCIes are new 0 scIence, an 0 er specIes,

previously known from outside Cambodia, represent new (C '\ F I ' b (16) Ann d 1 1 d thi .., . 1 d T . d.. f . d lore 1m s -mo erate y ong an n, J.orearmnatIona recor s. axonomIC ISCUSSIon 0 rare speCIes an(FLL 19 1 ) h h h d (HAL 20 0 )d .. f . 1. d .. d .mm sorter t an an .mm, notescnptIon 0 new specIes are Iste m systematIc or er. 1 d (17) F. 1 d th h. (TFL 10 9 )en arge .mgers ong an ra ert m .mm.(18) Relative length of fingers: II < I < IV < III. (19) Tips

fa' of all fingers rounded, slightly enlarged, without grooves.Lepto lax DuboIS, 1980 (20) Dennal fringe and webbing on fingers absent. (21)

." Subarticular tubercles indistinct, continuous ridge ofRemarks. -A .smgle specImen of thIS genus was collected thickened skin on underside of fingers. (22) Prepollex flat,from a mountam stream on Phnom Aural, 780 m a.m.s.l. It oval; palmar tubercles indistinct; supernumerary tuberclesis a larva at metamorphosis (stage 45) with its tail partially absent.regressed. It is rather large for its stage (SVL 26.1 mm), hasa granulose skin, distinctly smoother arms and elbows, and (D) Hind limbs -(23) Hind limbs long, heels overlappingits feet bear rudimentary webbing with fringes along the toes. when limbs are folded at right angles to body. Tibia fourVentral coloration in alcohol is unifonnly light grey without times longer (TL 38.8 mm) than wide (TW 10.6 mm), shorterdistinct spots. It is distinguished from the other species of than thigh (FL 39.2 mm), and longer than distance from basethis genus (Ohler et al., 2000) by its relatively long shanks of internal metatarsal tubercle to tip of toe IV (FOL 35.5(TL/SVL 598 p.m.). For specific detennination however, mm). (24) Toes long and thin, toe IV (FTL 17.9 mm) threestudy of adult specimens is necessary. This is the first record times the distance from base of tarsus to tip of toe IV (TFOLof this genus from Cambodia. 52.7 mm). (25) Relative length of toes: I < II < V < III <

IV. (26) Tips of all toes rounded, enlarged, without grooves.

(27) Webbing present, rudimentary, continued by a dennalMegophrys (Xenophrys) auralensis, new species fringe on toes: I -2 -21/2 II -2 -31/2 -III -3 -

(Fig. 1) 4 1/4 IV 4 1/4 -3 V (WTF n. m., WFF n. m., WI n. m.,WII n. m.; MTTF 11.3 mm, MTFF 12.6 mm, TFTF 20.0

Material examined. -Holotype -MNHN 2001.0209, adult male mm, FFTF 19.4 mm). (28) Dennal ridge along toe V absent.(Fig. I). Phnom [Mount] Aural in the Phnom Aural Wildlife (29) Subarticular tubercles indistinct, continuous ridge ofSanctuary, Kampong Speu Province, Southwest Cambodia (UTM thickened skin on underside of toes. (30) Inner metatarsal1326600N 0309200E). tubercle distinct and long, its length (IMT 5.83 mm) 1.2 times

the length of toe I (ITL 7.06 mm). (31) Tarsal fold absent.Diagnosis. -A large sized species of the subgenus Xenophrys (32) Outer metatarsal tubercle absent, supernumerarywith vomerine ridge, but without vomerine teeth, head rather tubercles absent; tarsal tubercle absent.broad, tympanum well developed, no white band on upperlip, tibia relatively long. (E) Skin -(33) Snout, between eyes, side of head, anterior

part of back smooth; posterior part of back and flanksDescription of holotype. -(A) Size and general aspect -(1) granular; palpebral horn indistinct. (34) Folds on back,Frog of large size (SVL 76.7 mm), body rather robust. including dorsolateral and middorsal folds outlining

hourglass pattern, narrow, fine. (35) Dorsal part of forelimb,(B) Head -(2) Head rather large, wider than long (HW 31.0 thigh, tibia and tarsus granular. (36) Throat, chest, bellymm; HL 28.0 mm, MN 23.4 mm; MFE 22.0 mm; MBE 13.5 and ventral part of thighs smooth. (37) Macroglands: smallmm), flat above. (3) Snout bluntly pointed, largely white femoral and pectoral glands; posterior part ofprotruding, its length (SL 7.6 mm) shorter than horizontal supratympanic fold scarcely enlarged.

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(F) Coloration -(In alcohol). -(38) Dorsal parts brown-grey with dark brown triangle between eyes and outlines ofdark brown hourglass pattern on back; flanks lighter greyishbrown with dark brown spots and whitish dots on largerglandular warts; canthus and tympanic fold dark brown;canthal and tympanic regions greyish brown; tympanum darkbrown; upper lip with dark vertical brown bands on greyishbrown ground colour; dorsal folds with brown outlines. (39)Forelimb, dorsal parts of thigh, tibia and foot greyish brownwith darker brown bands, posterior part of thigh uniformlydark brown. (40) Throat and chest dark brown; margin ofthroat with dark brown spots on white and brown speckledground colour; belly dark brown spots on whitish and brownspeckled ground colour; ventral part of thighs with densedark brown spots; webbing brown.

(In life) Dorsum dark tan with hourglass pattern and triangleon head dark mahogany brown; flanks indistinctly marbleddark brown; canthal and tympanic area darker brown, upperlip banded dark brown on vinous grey-brown ground colour.Dorsal surface of limbs and digits same colours as head andback: dark tan banded with darker brown; flushed orange ataxes of legs. Throat vinous grey-brown; belly and ventralsurface of limbs white with vinous grey-brown blotches;distinct pectoral glands white; posterior surface of thighs darkvinous brown with specks of white; webbing dark vinousbrown. Iris copper-coloured.

(G) Male secondary characters -(41) Nuptial pads on fingersI and II fonned by small, brown spines arranged in two ovalpads. (42) Vocal sacs present, indistinct on throat; a pair ofdistinct, rounded openings at base of jaw. (43) Othersecondary sexual characters absent.

Variation. -The nine males and the young female collectedare all very similar. There is some variation in the presenceof the dorsal hourglass pattern, which may be obscured bythe general coloration of the back. The holotype is lighterthan other specimens that are almost all of a dark browndorsal colour. The ventral coloration is very similar in patternand intensity in all specimens. A few show indistinct bandson the throat and chest, but never the clear-cut longitudinalshoulder stripes that can be observed in M. major.

Ecological notes. -The holotype was collected on PhnomAural at an altitude of 800 m a.m.s.l., calling from the topof a boulder 0.2 m from a cascade section of a mountainstream in hill evergreen forest. The collection data for theholotype are typical of this species on Phnom Aural: maleswere found to be common along mountain streams in hillevergreen forest from 500 to 1,140 m a.m.s.l. They wereseen and heard frequently calling from cascade sections ofthese streams. Typically found sitting upright on boulderswithin the splash zone of falling water, males were alwaysobserved within 1 m of the stream, and usually no furtherthan 0.3 m from fast-flowing water. Towards the end of thesurvey period, when the first "mango" rains had began tofall, male M. auralensis were heard to call during and afterrain throughout the day, and particularly at dusk. Suchcalling behaviour, i.e. from cascade sections of mountain

Fig. 1. Megophrys (Xenophrys) auralensis, new species. Ho1otype,MNHN 2001.0209, adult male, SVL 76.7 mm, dorsal view, ventralview, lateral view of head.

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Table 3. Minimum, maximum, mean and standard deviation of measurements and their comparison by Mann-Whitney U test of the typeseries of Megophrys auralensis, new species, and specimens of various origins of Megophrys major, including a syntype. (Initial SVLmeasurements are given in rom, all subsequent measurements are presented as per thousands of SVL). n = number; U = Mann-WhitneyU; p = probability; n.s. = not significant; * = significant; ** = very significant; *** = highly significant.

"IMeasurement M. auralensis n = 9 M. major n = 16 Mann-Whitney U test 1SVL 74.1 :I: 2.10 72.9:1: 5.88 U = 60.0 j

(71.0 -76.9) (63.7 -87.0) P = 0.522 n.s.

RHW 400 :I: 9.3 371 :I: 14.8 U = 5.0(383 -416) (346 -396) p = 0.000 ***

RHL 364 :I: 6.9 369 :I: 14.5 U = 55.0(352 -374) (339 -392) p = 0.357 n.s.

RFLL 254:1: 9.0 248:1: 11.9 U = 52.0(242 -271) (220 -262) p = 0.276 n.s.

RHAL 269:1: 8.9 249:t 12.2 U= 14.0(258 -287) (228 -276) P = 0.000 ***

RTL 522:1: 18.7 536:t 24.4 U = 47.0(491 -550) (488 -564) p = 0.169 n.s.

RFOL 473:t 16.6 508 :t 20.5 U = 14.0(442 -490) (466 -531) p = 0.000 ***

REL 127:1: 4.1 116:t 5.53 U = 8.0(120 -133) (106 -129) P = 0.000 ***

RTYD 81.1 :t 5.3 61 :t 10.6 U = 6.0(71 -91) (43 -81) p = 0.000 ***

RTYE 90.6:1: 6.3 80:t 4.7 U = 15.0(78 -96) (73 -90) P = 0.001 ***

RMTfF 155:t 13.7 177:t 23.8 U = 34.0(137 -176) (149 -215) P = 0.032 *

RMTFF 178:1: 11.9 204:1: 22.7 U = 21.0(161 -198) (168 -247) p = 0.003 **

RTFfF 272 :t 11.8 275 :t 22.5 U = 63.0(253 -285) (233 -304) p = 0.637 n.s.

RFFfF 262:t 12.3 275:t 17.1 U = 38.0(241 -276) (237 -302) p = 0.057 n.s.

streams, prompted by rain, is typical of other species of the head, and an absence of oblique dark bars, as are presentXenophrys (pers. obsv.). in M. auralensis. Biogeographically, the new species is

closest to M. major, but these two species can beEtymology. -The name auralensis, indicates that the species distinguished by several morphometrical characters (Tablewas found on Mount Aural, Cambodia's highest mountain. 3). Externally it can be recognized by the presence of dark

bands on the side of head, whereas in M. major there is aRemarks. -The specimens from Cambodia belong to a large distinct longitudinal whitish band on upper lip. Two Chinesesized form of Xenophrys (see Dubois & Ohler, 1988, for a species of this group, which show no band on upper lip,list of the species referred to this subgenus) showing should be compared to the new species. Megophrysrudimentary webbing on the hind feet. Between those omeimontis is of a smaller size (SVL 54.2-63.0 mm in thesharing these characters, M. gigantica Liu, Hu & Yang, 1960, five type-specimens described by Liu [1950]; U = 0.000, PM. glandulosa Fei, Ye & Huang, 1991, M. major (Boulenger, = 0.001) and has a longer tibia (521-605 p.t. SVL; U = 5.0,1908), M. mangshanensis Fei & Ye, 1991, and M. p=0.019). This species also has a smaller head (HW 339-shapingensis Liu, 1950, are distinct from the new species 375; U = 0.000, p = 0.001 / HL 322 -358; U = 2.0, P =by showing a well-defined white stripe on upper lip. In M. 0.004) than the new species from Cambodia, thus has to bemedogensis Fei, Ye & Huang, 1983, M. omeimontis Liu, considered specifically distinct. MegophrysmedogensisFei,1950, and M. major, vomerine teeth are present, whereas Ye and Huang, 1983 is morphologically very similar to M.such teeth are absent in specimens examined from the omeimontis, but has no webbing, whereas M. shapingensisCardamom,Mountains, which only show vomerine ridges lacks tympanum and has relatively more developed webbingdevoid of teeth. Megophrys robusta (Boulenger, 1908) continuing as a wide fringe on the toes.shows a larger head (412 p. t. SVL for the syntype measured,428 for a second specimen). It also can be distinguished by Inger (1999: 457) mentions "one single species ofthe presence of an indistinct unmarked area on the side of Megophrys" from the mountains in southeastern Thailand

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Table 4. Measurements (mm), means and standard deviations for specimens of Limnonectes gyldenstolpei from Thailand (including syntypes)and Laos, compared to specimens collected from the Cardamom Mountains in Cambodia.

Measurement Thailand and Laos including syntypes Cardamom Mountains, CambodiaMales N = 8 females N = 3 Males N = 3 Female

Snout-vent length 57.79:t 6.80 51.43 :t 2.57 63.0:t 9.06 54.4(49.7 -70.5) (49.9 -54.4) (52.6 -69.2)

Head width 26.53:t 3.48 20.20:t 1.20 29.0 :t 4.29 21.2(23.5 -33.0) (19.0 -21.4) (24.1 -31.9)

Head length 26.98 :t 3.93 20.77 :t 0.93 29.9 :t 4.08 22.9(23.3 -34.7) (20.0 -21.8) (25.2 -32.5)

Tibia length 28.76:t 3.18 26.13:t 0.25 31.5:t 4.10 28.5(23.0- 33.7) (25.9 -26.4) (26.8 -34.3)

-

and adjoining western Cambodia. In the summary table for Limnonectes (Elachyglossa) kohchangae (Smith, 1922)the region defined as Southeast Asian Lowlands, whichincludes central valleys of Myanmar and Thailand, Remarks. -Limnonectes kohchangae is considered to haveCambodia, southern and coastal Vietnam, and the Red River a restricted range being first described from Koh ChangValley of Vietnam, he then lists Megophrys longipes and Island in the Gulf of Thailand, and was only known fromM. parva. Concerning the presence M. longipes, we give this island and the islands of Koh Kut, Koh Mehsi, and fromsome remarks below. Whereas it is possible that M. parva Ok Yam on mainland Thailand (Smith, 1922a; Taylor, 1962).might be found in the Cardamom Mountains in the present However, whilst studying the material collected from thestudy area, it was not recorded during the present survey. Cardamom Mountains, two specimens (MNHN 1924.0065-

0066) of this species were discovered in the collection ofOnly one species of Xenophrys has been previously reported the Museum National d'Histoire Naturelle, Paris, donatedfrom Cambodia -Megophrys (Xenophrys) longipes by Smith in 1924, who collected them from Bokor in the(Boulenger, 1886). Tirant (1885) collected this species from Elephant Mountains of Southwest Cambodia, c. 150 kmthe Elephant Mountains, c. 100 km South of where the Sout~east of the Cardamom r~ge.. Bourret (1942: 265)Cardamom specimens were collected, but called it m~ntIons Ra~a koh~ha.ngae pouam from Dong Ta~ Ve,Megalophrys montana. Bourret (1942) cited these specimens Vlet~~m. Hl~ ?escnptI~n, ~as presented as a C~ndltIOnal !in the synonymy of Megophrys longipes in which he also one, 11 pourralt etre consldere comme appartenant a une race Iincluded a specimen from Vietnam, which has important differente', but the name is nevertheless available (Article I

h 1 . al dif" t th t t T .11.5.1 of Anonymous, 1999). The holotype from Vietnam,morp 0 OglC J.erences 0 e opo ypes. ype specImens ..f M I . f P k (M 1 . ) d .b d b figured by Bourret (1942: fig. 66) an adult male, IS m the0 .onglpes rom era a aysla escn e y 11 ' f h P . M (MNHN 19480127...co ectIons 0 t earls useum .ex

Boulenger (1886; 1908), have very long tIbIa (respectIvely LZNH B 298) d . d fi . 1 d " 1 .d (SVL.-an IS e Imte y a IstInct, arge sIze

574 and 585 p. t. SVL) and relatIvely small heads(HW 362 90 2 ) . fL '(El h I ) th h ldd 354 SVL) d h . f .mm speCIes 0 lmnonectes ac yg ossa at s ou

an .t. com are to t e s eclmens rom ...C b di Pth ld dp W al P d .bear the name Llmnonectes (Elachyglossa) pouam (Bourret,am 0 a at we cou stu y. e so compare specImens .

1942). Llmnonectes kohchangae has not been reported fromfigured by Bourret (1942) as Megophrys longipes (Table 3) Vietnam thus far. A Laotian record of Limnonectesand ~oncluded that ~ey could not ~e conspe~ific with M. kohchangae mentioned by Stuart (1999), citing a draft of alonglpes from MalaysIa. As th~ sp~clmens of Tlrant are lost, report to the Laotian authorities of Ohler (1997), is wrong.we cann?t confirm the determmatIon made. by Bourret: ~d Study of the Laotian voucher specimens indicate them beingM. l~nglpes should.be remo:ed from ~e lIst of amphIbIan of the Taylorana hascheana group (mentioned asspec~es of CambodIa an? VIetnam ~tIl we have v?ucher Limnonectes limborgi by Stuart 1999: 45). ComparisonspeCImens that confmn ItS presence m these countrIes. between the specimens collected during the present surveys

from the Cardamom Mountains, and the syntypes ofLimnonectes kohchangae from Natural History Museum

Limnonectes (Elachyglossa) gyldenstolpei (London), confirms that the specimens from Cambodia are(Andersson, 1916) conspecific with those from Thailand (Table 5).

Remarks. -This species was, until recently, known as Rana Limnonectes kohchangae was found to be widespreadpileata Boulenger, 1916, a name that has been shown to be throughout .the evergreen forests of the Cardamoma subjective junior synonym of Elachyglossa gyldenstolpei Mountains, from below 600 m a.m.s.l. to the top of the range,Andersson, 1916 (Ohler & Dubois, 1999). Our collection the highest record for this species being 1,200 m a.m.s.l.from the Cardamom Mountains includes three adult males, Often associated with forest streams but occasional, dispersedan adult female, and three juvenile specimens. Form and individuals were also recorded on the forest floor hundredssize of the dermal flap on the top of the head corresponds of meters from any body of water. In March of both years,to adult males from Thailand and Laos, as do major breeding choruses of L. kohchangae were observed besidemeasurements (Table 4). slow-flowing streams in upper hill evergreen forest.

471

,Ohler et al.: Amphibia of Cambodia

Table 5. Measurements (mm), means and standard deviations for specimens of Limnonectes kohchangae from Thailand (syntypes) comparedto specimens collected from the Cardamom Mountains in Cambodia.

--:4Measurements Thailand, syntypes Cardamom Mountains, Cambodia !

Male Female Males N = 8 Females N = 5

Snout-vent length 40.6 40.5 Co 42.0 :t 14.9 -~~.~-~ ~:84 (40.0 -44.0) (32.3 -40.8)

Head width 17.7 14.4 19.6:t 1.37 l5.4:t 1.95

(18.0 -21.0) (12.9 -17.1)Head length 17.8 15.2 19.4 :t 1.21 15.3 :t 1.85

(17.9 -20.9) (12.8 -16.7)Tibia length 19.9 20.3 20.8 :t 0.69 19.1 :t 1.98

.(19.4 -21.7) (15.7 -20.7)

Table 6. Measurements (mm) of type specimens of Faa jasciculispina, as published in Inger (1970) of topotypic collection, and materialcollected from the Cardamom Mountains in Cambodia.

, '~.~~ .Measurement Thailand

Holotype Paratype Topotypes males Topotype femaleN=3 'I;-.-

Snout-vent length 106 104 108.37 :t 5.51 101.1

Head width 43 41 43.27:t 1.95 39.1

Head length 42 40 41.03:t 1.40 39.3

Tibia length 53 49 53.93 :t 4.40 49.3

Cardamom Mountains, CambodiaTopotype young Adult male Adult female Young

N=5Snout-vent length 94.3 100.9 108.3 47.74 :t 8.38

Head width 37.7 40.3 45.2 19.06:t 3.27

Head length 35.7 39.3 41.1 18.6:t 3.05

Tibia length 46.7 47.6 49.5 23.3 :t 4.07

As a consequence of an apparent restricted range L. The populations of this species from the Cardamom

kohchangae is listed as "Threatened" by the IUCN (2001), Mountains are without doubt con specific with those inbut under the Data Deficient category, The conservation Thailand,

status of this species should be reviewed in the light of the

present discovery of L. kohchangae in the Cardamom PaafascicuZispina was recorded in streams flowing throughMountains. hill evergreen forest, at altitudes of c. 700-1,000 m a.m.s.l,

During March, male P. fascicuZispina were heard calling

from concealed positions, under boulders, in cascade sectionsPaa (Eripaa) fasciculispina (Inger, 1970) of mountain streams on Mount Aural and Mount Samkos,

Remarks. -The specimens of Paa fascicuZispina collected

in the Cardamom Mountains represent the first record of thePh . l t .l .

'. z au us caruamonus, new species

ranld genus Paa from Cambodia and the second record of (F. 2)this species since its description by Inger in 1970. Ig. 1

Considering the type locality (in Chanthaburi Province, , .S th t Th .1 d) f th ' ' p fi . Z '. Id Matenal examzned. -Holotype, BMNH 2000.0149, adult male,ou eas at an 0 IS specIes, .asczcu zspma wou. ...b t d t . h b 't th " t tr f th C d (FIg. 2). Phnom Sarnkos ill the Phnom Sarnkos WIldlIfe Sanctuary,e expec e 0 ill a I e lores seams 0 e ar amomM nw ' Pursat Province, Southwest Cambodia (UTM 1343841 N, 0287890

ou ns,E).

Morphological comparison (Table 6) to type specimens '. '. ..i(Inger, 1970) and to topotypes from the Paris collection Dzagnoszs. -Small sized Phzlautus Without vomenne teeth; J

reveals a close similarity between the Cambodian and Thai nuptial pad present; m. cutaneus pectoris absent; m.

specimens, and secondary sexual characters of Cambodian geniohyoideus medialis free; ova half pigmented; feet half

males are consistent with the description of the holotype. webbed; ventral body with distinct dark and light marbling.

472


Description of holotype. -(A) Size and general aspect -(1)Frog of small size (SVL -19.3 mm), body rather stout.

(B) Head -(2) Head moderate size, longer than wide (HW7.3 mm; HL 8.2 mm, MN 7.37 mm; MFE 6.18 mm; MBE2.89 mm), convex above. (3) Snout rounded, slightlyprotruding, its length (SL 3.44 mm) equal to horizontaldiameter of eye (EL 3.44 mm). (4) Canthus rostralis rounded,loreal region concave, flared. (5) Interorbital space convex,larger (IUE 3.05 mm) than upper eyelid (UEW 2.14 mm)and than intemarial distance (IN 2.27 mm); distance betweenfront of eyes (IFE 4.86 mm) about two times in distancebetween back of eyes (IBE 8.43 mm). (6) Nostrils roundedwithout flap of skin laterally, slightly closer to tip of snout(NS 1.62 mm) than to eye (EN 1.81 mm). (7) Pupil oval,horizontal. (8) Tympanum (TYD 1.43 mm) distinct, rounded,42% of eye diameter; tympanum-eye distance (TYE 0.39mm) 27% of tympanum diameter. (9) Pineal ocellus absent.(10) Vomerine ridge absent. (11) Tongue moderately,cordate, emarginate, bearing no median lingual process.Tooth-like projections on lower jaw absent. (12)Supratympanic fold distinct, from back of eye to aboveshoulder. (13) Parotoid glands absent. (14) Cephalic ridgesabsent. (15) Co-ossified skin on head absent.

(C) Forelimbs -(16) Ann short and thin; forearm (FLL 5.25mm) shorter than hand (HAL 7.26 mm), not enlarged. (17)Fingers I and II short and thin; fingers III (TFL 4.21 mm)and IV long and thin. (18) Relative length of fingers: I < II< IV < III. (19) Tips of all fingers with well-developed disks,with distinct circummarginal grooves, rather wide comparedto finger width (fdl 0.65, fwl 0.43 mm; fd2 0.90 mm, fw20.43 mm; fd3 1.13 mm, fw3 0.85 mm; fd4 1.10 mm, fw40.43 mm). (20) Dermal fringe on inside of fingers I to III;webbing on fingers absent. (21) Subarticular tuberclesdistinct, rounded, single, all present. (22) Prepollex distinct,oval; two palmar tubercles; supernumerary tubercles presenton all fingers.

(D) Hind limbs -(23) Hind limbs long, heels overlappingwhen limbs are folded at right angles to body. Tibia fivetimes longer (TL 11.1 mm) than wide (TW 1.9 mm), longerthan thigh (FL 10.2 mm) and distance from base of internalmetatarsal tubercle to tip of toe IV (FOL 10.66 mm). (24)Toes moderately long and thin, toe IV (FTL 5.70 mm) 2.5times the distance from base of tarsus to tip of toe IV (TFOL14.5 mm). (25) Relative length of toes: I < II < III < V < IV.(26) Tips of all toes with moderate disks, with distinctcircummarginal grooves, moderately wide compared to toewidth (td1 0.59 mm, tw1 0.50 mm; td2 0.70 mm, tw2 0.47mm; td3 0.74 mm, tw3 0.50 mm; td4 0.90 mm, tw4 0.54mm; td5 0.95 mm, tw5 0.65 mm). (27) Webbing present,moderate: I -2 -'-- 21/2 II -1 -21/2 -III -11/2 -21/2 IV 21/2 -11/2 V (WTF 1.34 mm, WFF 1.80 mm, WI1.03 mm, WII 1.24 mm; MTTF 4.99 mm, MTFF 5.64 mm,TFTF 4.99 mm, FFTF 4.41 mm). (28) Dermal ridge alongtoe V present, from tip of toe to proximal subarticulartubercle. (29) Subarticular tubercles prominent, rounded,simple, all present. (30) Inner metatarsal tubercle distinctand short, its length (IMT 0.90 mm) 2.9 times the length of

Fig. 2. Philautus cardamonus, new species. Holotype, BMNH2000.0149, adult male, SVL 19.3 mm, dorsal view, ventral view,lateral view of head.

473


toe I (ITL 2.58 mm). (31) Tarsal fold absent. (32) Outer wildlife sanctuary. These frogs were also found on leavesmetatarsal tubercle absent, supernumerary tubercles present of plants approximately 2.5 m from a mountain stream.on all toes; tarsal tubercle absent.

Etymology. -Cardamonus, adjective, derived from the Latin(E) Skin -(33) Snout, between eyes, side of head, anterior "cardamonus", the name of the plant esteemed for its seedsand posterior part of back and flanks shagreened. (34) Dorso- that are used as a spice in cooking and in traditional medicine.lateral folds absent. (35) Dorsal part of forelimb, thigh, tibia The specific name refers to the type locality in the Cardamomand tarsus shagreened. (36) Throat, chest, belly and ventral Mountains.part of thighs granular ("tree frog belly skin"). (37)Macroglands: small white rictal gland posterior to comer of Remarks. -Character combination in Philautus cardamonusmouth. does not fit with any of species groups as defined by Dring

(1987). It shares with species from the vermiculatus group,(F) Coloration -(In alcohol). -(38) Dorsal parts bluish grey the absence of m. cutaneus pectoris and the presence of awith dark grey brown pattern, notably a large triangular spot free geniohoideus medialis and the pigmented ova. Howeverbetween eyes separating into two bands continuing laterally the males of the vermiculatus group are devoid of nuptialto groin; lower flanks cloudy dark brown; canthus and pads, whereas the holotype bears distinct pads on first fingertympanic fold dark brown; canthal and tympanic area and prepollex. The dorsal pattern is very similar to P.greyish; tympanum light grey; upper lip with snow-white carinensis (Boulenger, 1893) and P. jinxiuensis Hu, 1978.spots on greyish ground. (39) Forelimb, dorsal parts of thigh, However both these species do not have a distinct ventraltibia and foot fawn with some darker brown bands, posterior pattern, even if some individuals of P. carinensis show apart of thigh yellow-orange. (40) Throat, margin of throat greyish shade on the vent. The size of adult males in bothand chest yellow white with very indistinct brown marbling; species is superior to the size of the holotype (23.2 -33.9belly yellow white with round white spots; ventral part of mm in six adult males of P. carinensis from Sa Pa, Vietnam;thighs yellow white; webbing greyish. 23.5 mm in P. jinxiuensis according to Fei et al. [1999]).

The presence of a distinct tympanum distinguishes this(In life) Dorsum dark tan with a pattern of dark mahogany species from species such as P. banaensis Bourret, 1939, P.brown; flanks indistinctly marb(ed dark brown; canthal and gryllus Smith, 1924, P. parvulus (Boulenger, 1893), P.tympanic area same dark brown. Dorsal surface of limbs kempiae (Boulenger, 1919) and P. annandalii (Boulenger,same colours as head and back: dark tan banded with darker 1906). The webbing of P. cardamonus is somewhatbrown. Throat and chest white marbled with dark brown intermediate and less developed than in P. carinensis" P.blotches; belly and ventral surface of limbs greenish yellow. maosonensis and P. banaensis, but more developed than P.Iris copper-coloured. parvulus, P. gryllus and P. annandalii, which have only basic

webbing on the feet. The presence of distinct white spots(G) Male secondary characters -(41) Nuptial pads present, on the flank easily distinguishes this species of othersingle oval patch from base of finger I to level of subarticular Philautus.tubercle, indistinct ivory white spines. (42) Vocal sacspresent, indistinct on throat; a pair of distinct, slit openings The first record of this genus is from Smith (1930) whoat base of jaw. (43) Other secondary sexual character: not mentioned Northern Siam (Khum Tan) and Southernobserved. Cambodia as the northern limits of the range of Philautus

petersi. This information was also repeated by BourretVariation. -The two female paratypes show the same dorsal (1942). Smith (1930) does not cite voucher specimens, sopattern as the male, but ventrally and laterally they show a specific allocation cannot be confirmed. Dring (1987)very distinct white and dark brown vermiculation, less consigned Philautus petersi to the Philautus aurifasciatusdistinct on throat and lower belly. In the male this pattern group. This might indicate that the specimens mentioned byis indistinct, visible only from the presence of the white Smith (1930) are not conspecific with our sample. Philautusrounded spots on the belly. In life the paratypes are of a petersi has not been mentioned from Thailand and Cambodialighter colour than the holotype: the dorsal surfaces of both in more recent works (Taylor, 1962; Frost, 2000) nor couldbody and limbs being a very pale gold colour with patterns we find mention of Smith's specimens in Taylor (1962).of copper rather than dark brown. The venter is white witha more distinct dark grey-brown marbling that extends on tothe flanks and upper lip; the ventral surfaces of the limbs Rana (Sylvirana) Dubois, 1992are yellow. Iris colour of the paratypes in life is an iridescentmix of gold and copper. Remarks. -Two groups of ranid frog belonging to the sub-

genus Sylvirana were collected from the CardamomEcological notes. -The holotype was collected on Phnom Mountains. These two frogs are from the Rana nigrovittataSarnkos, in upper montane evergreen forest at an altitude of complex. One species should be given the name Rana1,650 m a.m.s.l. This specimen was found on the leaf of a mortenseni Boulenger, 1903. We compared the Cambodianbush, hundreds of meters from any source of water. The specimens to a syntype and a topotype of Rana mortenseniparatypes were collected in a similar forest type at the lower from Koh Chang Island, but also to a series of specimensaltitude of 1,250 m a.m.s.l. on Phnom Tumpor in the same from northeastern Thailand, that all share the short-snouted,

I474


large head and relatively large body size of Rana mortenseni axis, as close to choanae as to each other, longer than distance(see Table 7 for measurements). The validity of this species between them. (11) Tongue moderate, spatulate, emarginate,has been confirmed by comparison with the lectotype and bearing no median lingual process. Tooth-like projectionsparalectotypes of Rana nigrovittata (Blyth, 1855). Matsui on lower jaw absent. (12) Supratympanic fold absent. (13)et al. (2001) found at least two genetically distinct subgroups Parotoid glands absent. (14) Cephalic ridges absent. (15)(Nei distance> 0.2) in their sample of the Rana nigrovittata Co-ossified skin on head absent.group from Thailand. This genetic differentiation isstrengthened by morphological differentiation (Ohler, (C) Forelimbs -(16) Ann moderate; forearm (FLL 13.8 mm)unpublished data) but as many as five different taxa might shorter than hand (HAL 15.9 mm), slightly enlarged. (17)be recognized in Thailand and the Indochinese subregion Finger II short and thin; fingers I, III (TFL 9.3 mm) and IVfor this species group (Ohler, unpublished data). long and thin. (18) Relative length of fingers: II < I < IV

< III. (19) Tips of all fingers pointed with disks, with latero-The second species found on the Cardamom Mountains is ventral grooves, moderately wide compared to finger widthdistinct from all known species of Sylvirana. It is a large (fdl 1.37 mm, fwl 0.87 mm; fd2 1.24 mm, fw2 1.09 mm;sized form, reminding Rana.8uentheri Boulenger, 188~,"but fd3 1.34 mm, fw3 0.87 mm; fd4 1.40 mm, fw4 1.00 mm).males of the Cardamo~ s~ecimens have no ext~rna1ly vIsIble (20) Dermal fringe on inside of fingers absent; webbing onvocal sacs a~d are dIS!mctly smaller. ThIS t.axon. was fingers absent. (21) Subarticular tubercles very prominent,separate~ earlIer by SmIth (1922b) from R.ana mgr~vltt~ta oval, single, all present. (22) Prepollex distinct, oval; twosensu stricto and also from Rana mortensem, but considenng oval distinct palmar tubercles. a distinct supernumeraryit a variation, he did not name it. We will formally name tube;cle on base of each finger:the second species from Cardamom Mountains in memoryof the work of Malcolm Smith on Southeast Asian (D) H . d l . b (23) H . d 1. b 1 h 1 1 . m lm s -m 1m song, ee s over appmg

amphIbIans, partIcularly on some dIfficult groups such as h 1. b fi Id d t . ht 1 t b d T ' b ' .w en 1m s are 0 e a ng ang es 0 0 y. 1 Ia more

Llmnonectes (Elachyglossa) and Rana (Sylvirana). th fi ti. 1 (TL 37 9 ) th .d (TW 8 1an our mes onger .mm an WI e .

mm), longer than thigh (FL 33.6 mm) and distance from baseR (S l . ) fi b .of internal metatarsal tubercle to tip of toe IV (FOL 35.2

ana y Vlrana a er, new speCIes ) (24) T 1 d h. IV (FfL 20 8 ) 2 5(F. 3) mm .oes ong an t m, toe .mm .

Ig. times the distance from base of tarsus to tip of toe IV (TFOL

0" t . l . d H 1 MNHN 20010261 d 1 1 50.6 mm). (25) Relative length of toes: I < II < III < V <",~a ena examlne .-0 otype, "' aut ma e IV (26) T. f all(F.

3) Ph A 1. th Ph A 1 W " ldl.f S .IpS 0 toes poInted WIth small dISks possessIng19. .nom ura III e nom ura lie anctuary, ..

K S P . S h C b d " (UTM 1328200N latero-ventral grooves, rather wIde compared to toe wIdthampong peu rovillce out west am 0 la

0307700E). (tdl 1.62 mm, twl 0.87 mm; td2 1.71 mm, tw2 1.09 mm;td3 1.71 mm, tw3 0.90 mm; td4 1.49 mm, tw4 0.93 mm; td5

Diagnosis. -Large sized Sylvirana with relatively narrow 1.56 mm, tv:5 0.87 mm). (27) Webbing present, rather large:head, vocal sacs externally not visible, humeral gland 1-0 -1/211 -0 -2 -111- 0 -2 IV 2 -0 V (WTFindistinct. Dorsal skin finely granular forming horny spinules 9.61 mm, WFF 9.34 mm, WI 8.82 mm, WII 7.63 mm; MTfFon posterior back, narrow prominent dorsolateral folds. 19.4 mm, M~ 20.9 mm, TFTF 12.9 mm, FFT~ 13.0 mm).Dorsal colour light tan, limbs paler with indistinct bands, (28) Dermal ndge along toe V present, from tip of toe tothroat s~e colour than chest and belly. external metatarsal tubercle. (29) Subarticular tubercles

prominent, oval, simple, all present. (30) Inner metatarsalDescription ofholotype. -(A) Size and general aspect -(1) tubercle distinct and rather short, its length (IMT 3.16 mm)Frog of rather large size (SVL 59.4 mm), body elongate. 2.8 times the length of toe I (ITL 8.95 mm). (31) Tarsal

fold absent. (32) Outer metatarsal tubercle rounded,(B) Head -(2) Head moderate size, longer than wide (HW prominent; supernumerary tubercles absent; tarsal tubercle19.8mm;HL22.8mm,MN 19.7 mm; MFE 13.7 mm;MBE absent.

7.2 mm), flat above. (3) Snout rather pointed, slightlyprotruding, its length (SL 10.00 mm) longer than horizontal (E) Skin -(33) Snout, between eyes, side of head and anteriordiameter of eye (EL 7.89 mm). (4) Canthus rostralis rounded, part of back finely granular; posterior part of back and upperloreal region concave, vertical. (5) Interorbital space flat, part of flanks with small glandular warts bearing hornynarrower (IUE 5.13 mm) than upper eyelid (UEW 6.18 mm) spinules; lower part of flanks rather smooth. (34) Dorso-and than internarial distance (IN 6.05 mm); distance between lateral folds prominent, rather narrow. (35) Dorsal part offront of eyes (IFE 11.1 mm) two thirds of distance between forelimb with small glandular warts; thigh, tibia and tarsusback of eyes (IBE 15.7 mm). (6) Nostrils oval with flap of with small glandular warts in longitudinal lines bearing hornyskin laterally, closer to tip of snout (NS 4.34 mm) than to spinules. (36) Throat, chest, belly and ventral part of thighseye (EN 5.26 mm). (7) Pupil oval, horizontal. (8) smooth; posterior part of thigh around vent with denseTympanum (TYD 5.39 mm) distinct, rounded, 70% of eye glandular warts. (37) Macroglands: distinct rictal glanddiameter; tympanum-eye distance (TYE 1.97 mm) 37% of posterior to comer of mouth; flat indistinct humeral glands.tympanum diameter. (9) Pineal ocellus present, betweenanterior border of eye. (10) Vomerine ridge present, bearing (F) Coloration -(In alcohol). -(38) Dorsal parts light grey-numerous (N=8) small teeth; with an angle of 450 to body brown; dark grey stripe below the dorsolateral folds, lower

475 ,II


flanks light grey with blackish speckles; canthus dark grey;canthal and tympanic area light grey; tympanum brownishgrey, transparent; upper lip with pearl-white stripe continuedby pearl-white rictal gland; dorsolateral fold medially greyishbrown, externally black underlining. (39) Forelimb, dorsalparts of thigh, tibia and foot greyish brown with dark greybands, posterior part of thigh blackish with numerous lightgrey spots. (40) Throat and margin of throat grey-white withgreyish flecks; chest light grey with two symmetrical greymarks in the middle; belly cream-white with indistinctgreyish spots; ventral part of thighs yellow-white withgreyish spots; webbing dark grey, transparent.

(In life) Dorsal ground colour uniform light tan; flanks darkbrown; canthal and tympanic area dark brown; tympanumdark brown; upper lip pearly white continuous with pearlywhite rictal gland. Dorsal surfaces of limbs paler than backwith indistinct darker brown bands; posterior part of thighblotched dark brown on light tan ground colour. Venterpearly white; limbs flushed pink.

(G) Male secondary characters -(41) A single continuous,oval shaped nuptial pad on finger I formed by indistinct,small, cream coloured spines. (42) Vocal sacs present,indistinct on throat; a pair of distinct, rounded openings atbase of jaw. (43) Other secondary sexual characters: forearmslightly enlarged; humeral gland present but little prominent.

Variation. -Morphometric variation is presented in table 7.Dorsal colour in most specimens is greyish brown, but manyshow light grey or blackish flecking reminding lichens. Thethroat never shows distinct darker coloration than chest andbelly. Vocal sacs are indistinct externally; sometimes a darkerzone can be observed where the vocal sacs are located. Thehumeral gland, that shows thickened glandular tissue whendissected, is indistinct by superficial observation in all males.

Ecological notes. -The holotype was collected from PhnomAural in hill evergreen forest at an altitude of 710 m a.m.s.l.The specimen was found sitting on a rock. 0.4 m from astream. Rana Jaber was found to be common and widespreadthroughout the Cardamom Mountains occurring in a rangeof vegetation types: dry dipterocarp. lowland dry evergreenand hill evergreen forest types, moorland areas on the top ofthe central range, and flooded forest in the Veal Vengwetland. Rana Jaber was recorded in heavily disturbed areasas well as in near-pristine forests. Typically this specieswas found close to permanent watercourses, often slowermoving sections of mountain streams.

Etymology. -This species is dedicated to Malcolm A. Smithin recognition of his work on Southeast Asian amphibians.Faber in Latin means "smith" or "craftsman working hardmaterial".

Remarks. -Rana (Sylvirana)faber is compared here to twosimilar species, R. nigrovittata and R. mortenseni. AlthoughR. mortenseni was recently regarded as a junior subjectivesynonym of Rana nigrovittata by Matsui et al. (2001), wehave unpublished infonnation indicating that the two species

Fig. 3. Rana (Sylvirana) faber, new species. Holotype, MNHN2001.0261, adult male, SVL 76.7 mm, dorsal view, ventral view,lateral view of head.

476


Table 7. Minimum, maximum, mean and standard deviation of measurements of specimens of Rana (Sylvirana) mortenseni from Thailandand the Cardamom Mountains, Cambodia, and the type series of Rana (Sylvirana) faber, new species, from the Cardamom Mountains,Cambodia. (Initial SVL measurements are given in mm, all subsequent measurements are presented as per thousands of SVL).

Rana mortenseni Rana faber, new speciesThailand Cardamom Mountains, Cambodia Cardamom Mountains, Cambodia

adult male adult female adult male adult female adult male juvenile male adult female! N=ll N=6 N=8 N=2 N=9 N=2 N=3

~ Snout-vent 64.5 ::!: 6.37 67.8::!: 2.38 65.4::!: 6.57 63.8 -79.8 60.0::!: 3.72 38.5 -41.1 57.4::!: 0.35

length (54.3 -72.3) (64.6 -70.7) (51.2 -73.4) (53.1 -66.9) (57.1 -57.8)

Head width 362::!: 19.4 343::!: 14.1 380::!: 7.72 314 -327 321::!: 10.7 316 -343 344::!: 10.5ratio (319 -387) (322 -356) (367 -392) (306 -339) (336 -356)

Head length 395::!: 10.8 382::!: 6.29 414::!: 8.24 370 -389 381 ::!: 10.5 364 -384 387::!: 17.8ratio (374 -415) (375 -388) (405 -431) (365 -397) (367 -401)

Tibia length 527 ::!: 26.6 540::!: 13.7 535 ::!: 21.3 618 -630 618 ::!: 16.5 569 -600 558 ::!: 19.2ratio (488 -567) (515- 552) (500 -557) (594 -638) (536 -571)

are indeed distinct. Therefore, for the time being and for chaseni Smith, 1924 from the Malay Peninsula. All butpurposes of comparison, R. nigrovittata and R. mortenseni verrucosus are considered synonyms of Rh. appendiculatusare regarded here as separate species. (Frost, 2000).

Rana (Sylvirana) Jaber is distinctly larger than Rana More recently, Taylor (1962) described Rhacophorusnigrovittata, it also has longer tibia than this species. In R. bisacculus without comparison to the other species of thisnigrovittata, vocal sacs are distinct on a throat that generally group. Fei & Ye (Ye et al., 1993) coined the name Philautusshows a dark brown coloration in males. Rana mortenseni odontotarsus for the Chinese specimens considered until thenis a little larger size but can easily be recognized by its as Rhacophorus appendiculatus, and Inger et al. (1999)enlarged short-snouted head, and the shanks of R. mortenseni named the southern Vietnamese population Rhacophorusare significantly shorter than those of R. Jaber. baliogaster.Morphometric measurements of specimens of R. mortenseniform the Cardamom Mountains, plus a series from Thailand, Here, we follow Inger et al. (1999) in considering Rh.and those of the type series of R. Jaber are presented for appendiculatus a species distinct from Rh. verrucosus, andcomparison in Table 7. The dorsal pattern of R. mortenseni Rh. bisacculus different again from both of these, based onis also a distinctive reddish brown, and the chest and throat size and differences in body proportions. The specimensare dark brown (in some specimens even the belly is dark from the Cardamom Mountains possess body and tympanumbrown). Rana mortenseni also shows a rounded, well- sizes corresponding to those measured for Rhacophorusdeveloped humeral gland. This species pair can also be bisacculus. Van Dijk (unpublished data) mentions Rh.clearly separated in the field by their distinct calls, in addition verrucosus from Cambodia, but as no voucher specimensto their morphological differences and habitat preferences. are available, these specimens should be considered Rh.The Cardamom members of the subgenus Sylvirana exhibit bisacculus.some degree of altitudinal zonation: R. mortenseni isabundant in the lowland dry evergreen and gallery forestsof the basin areas and lower slopes but was rarely found DISCUSSIONabove c. 700 m a.m.s1.; R. Jaber, although found at loweraltitudes, was more common on streams above the extent of In terms of species composition of the Cardamom Mountain

f R. mortenseni's ecological range. amphibi~ fauna, the results of these s~r:eys have producedfew surpnses. The lowland COmmUnIties of the open drydipterocarp forests, at the base of the mountains, are

Rhacophorus bisacculus Taylor, 1962 comprised of common species widely distributed throughoutSoutheast Asia, that often characterise disturbed or

Remarks. -The oldest available name for small-sized agricultural habitats. Of the species of the evergreen forestsrhacophorids from the oriental region with an hourglass of the mountains themselves, although many never recordedpattern on the back and tubercles on the margins of the tarsi in Cambodia before, most have been reported from otherand feet is Rhacophorus appendiculatus (Giinther, 1859), parts of the Indochinese region, particularly from easternwhich was described from a variety of type localities Thailand. The data presented here merely fills in the gapsincluding the Philippines, Java, Singapore and the East (or confirms predictions) in the distribution of manyIndies. Subsequently, Rhacophorus verrucosus Boulenger, .Indochinese amphibians.1893 was described from Burma; Rhacophorus phyllopygus .Werner, 1900 from Sumatra; Rhacophorus naso Annandale, Despite the relative geographical isolation of the Cardamom1912 from Arunachal Pradesh, India; and Rhacophorus Mountains, their restricted altitude combined with a limited

477


temporal isolation has not been sufficient to produce The incompleteness of data sets, and the variable levels ofparticularly high levels of endemism. Three species of survey effort for sites already studied, hinders validamphibian new to science from the Cardamom Mountains comparisons of species richness between sites. Despite thesecollection could be considered potentially endemic to these restrictions, it is apparent that the many and varied habitatsmountains: Megophrys auralensis, Philautus cardamonus, that comprise the Cardamom Mountain area support aand Ranafaber. Philautus is the most speciose of frog genera relatively rich amphibian fauna, including a few taxa whichin the Orient, characterised by direct development (Bossuyt have evolved in isolation to produce species endemic to the& Dubois, 2001) and concomitant restricted specific mountain range.distribution and high levels of endemism. It is not uncommonfor surveys of previously unstudied forests in Indochina to Forests in Cambodia were once, ironically, protected to someyield at least one new species of Philautus, e.g. Inger et al. degree by an ongoing civil war. In the last few years, since(1999), Ohler et al. (2000) and Swan (unpublished data). attaining relative political stability, the forests of theTherefore, it was not unexpected to find at least one new Cardamom Mountains have become vulnerable to bothspecies from the higher elevations of the Cardamom human encroachment and commercial logging. At the timeMountains. Further searching at the higher elevations is of the survey, development aid agencies were activelybound to yield more species, rare and potentially endemic, resettling refugees within the two protected areas in thewhich are inevitably the last to be discovered. Cardamom Mountains, while between the protected areas,

the forest had been placed under concession to a number ofEndemicity at the species level in the Cardamom Mountains commercial logging companies.appears to be somewhat restricted in other vertebrate groups.For example, only a single species of terrapin (Cyclemys This preliminary inventory of the amphibian species of theatripons Iverson & McCord, 1997), lizard (Cyrtodactylus Cardamom Mountains was part of a larger effort to documentintermedius Smith, 1917), snake (Lycodon sp.) and bird and conserve the fauna and flora of this vast forested(Arborophila cambodiana Delacour & Jabouille, 1928) seem wilderness. Until the 2000 dry season surveys of amphibiansto be endemic to the mountains of Southwest Cambodia and and other taxa, both wildlife sanctuaries lacked activeadjacent eastern Thailand to date (Daltry & Chheang pany, management on the ground. Results of the biological surveys2000; Daltry & Wiister in press; Steinheimer et al., 2000), have provided the justification to establish the fIrst rangeralthough further research may find more. In the case of all forces in these mountains and begin developing managementof these examples, and that of the two new species of frog plans for the protected areas, while logging concessions indescribed here, the Cardamom endemic species are very the Central Cardamoms are being revoked now that thesimilar in external morphology to closely related congeners biological value of these forests is becoming apparent. It isfrom other parts of Southeast Asia. This suggests that the hoped that these initial steps towards enforced protection offauna of the Cardamom Mountains has undergone only the Cardamom Mountains will constitute the foundations oflimited divergent evolution in isolation. an UNESCO World Heritage Site nomination, a prestigious

profile that is necessary to safeguard the future of one of theLittle has been recorded of Cambodia's amphibian fauna largest and most intact areas of forest cover in mainlandprior to this study and a synthesis of this information has Southeast Asia.never been produced, thus limiting comparisons of overallspecies richness between the results presented here with those The survey of the Cardamom Mountains reported herein,from other parts of the country. However, the number of was the first survey for amphibians to be conducted inspecies recorded in the Cardamom Mountains (34 in total) Cambodia. Following immediately after the 2000 surveysis higher than known levels of species richness documented of in the Cardamom Mountains, the first author conductedfor other Indochinese mountainous areas of comparable a rapid survey of the plains of Northeast Mondulkiri onelevation. For example, Robichaud & Stuart (1999) opposite side of the country, close to the border with Dakdocumented 25 species of amphibian during a one-month Lak Province in Vietnam. This low-lying «200 m) plain atsurvey in the Annamite Mountains in Laos. Similarly, at a the foot of the Chhlong Plateau is part of the lower Mekongsite on the Vietnamese side of the same mountain chain, 30 basin and is clad in dry dipterocarp forest (Long et al., 2000)species of amphibian were recorded during a four-month much the same as that found in the basin areas at the footsurvey (Swan unpublished data). Yet the Cardamom of the Cardamom Mountains. Only 14 species of amphibianMountain amphibian fauna appears not to be as species rich were recorded during this survey, all but one, Ranaas the mountain isolates of the region that attain altitudes in (Pelophylax) lateralis Boulenger, 1887 (BMNH 2000.0130-excess of 2,500 m a.m.s.l., such as the Central Highlands of 0132), of which were recorded in the Cardamom Mountains.Vietnam (54 species -Inger et al., 1999; Le Trong Trai et The inventories produced by these two preliminary studies,al., 1999, 2000; Tordoff et al., 2000), or the Hoang Lien coupled with a few other incidental reports comprise theMountains, again in Vietnam (42 species -Ohler et al., 2000). known amphibian fauna of Cambodia. A checklist of the

fauna is presented in Table 8, which lists a total of 40 speciesNone of these examples, or the surveys reported herein, can of amphibian recorded from Cambodia.be considered complete. There are undoubtedly more speciesof amphibian to be discovered throughout the mountainous Surveys for amphibians in other areas of Cambodia (notablyregions of Indochina, including the Cardamom Mountains. the Elephant Mountains and southern Mondulkiri Province)

478

THE RAFFLES BULLETIN OF ZOOLOGY 2QO2

Table 8. Checklist of amphibian species known to occur in Cambodia.

SPECIES REFERENCE

MEGOPHRYIDAE Bonaparte, 1850

Leptolalax sp. Present studyMegophrys (Xenophrys) auralensis, new species Present study

BUFONIDAE Gray, 1825

Bufo galeatus Gunther,1864 Gunther, 1864; van Dijk unpublished dataBufo macrotis Boulenger, 1887 Long et al., 2000; this paperBufo melanostictus Schneider, 1799 Flower, 1896; Mocquard, 1904; Long et al., 2000; van Dijk

unpublished data; present studyBufo parvus Boulenger, 1887 Present study

RANIDAE Rafinesque-Schmaltz, 1814

Chirixalus doriae Boulenger, 1893 Present studyChirixalus vittatus (Boulenger, 1887) Present studyFejervarya cancrivora (Gravenhorst, 1829) Bourret, 1942 ;Fejervarya limnocharis (Gravenhorst, 1829) Long et al., 2000; van Dijk unpublished data; present study iHoplobatrachus chinensis (Osbeck, 1765) Long et al., 2000; present studyLimnonectes (Elachyglossa) gyldenstolpei (Anderson, 1916) Present studyLimnonectes (Elachyglossa) kohchqngae (Smith, 1922) Present studyLimnonectes toumanoffi (Bourret, 1941) Bourret, 1941; Ohler & Dubois, 1999; van Dijk unpublished dataOccidozyga lima (Gravenhorst, 1829) Boulenger, 1882; Flower, 1896; Bourret, 1942; Long et al.,

2000; van Dijk unpublished data; present studyPaa (Eripaa) fasciculispina (Inger, 1970) Present study iPhilautus cardamonus, new species Present study !Philautus parvulus (Boulenger, 1893) Present studyPhrynoglossus martensii Peters, 1867 Flower, 1896; Long et al., 2000; van Dijk unpublished data;

Present studyPolypedates cf. leucomystax (Gravenhorst, 1829) Long et al., 2000; van Dijk unpublished data; present studyRana (Hylarana) erythraea (Schlegel, 1837) Tirant, 1885; van Dijk unpublished data; present studyRana (Hylarana) macrodactyla (Gunther, 1858) Boulenger, 1920; Bourret, 1942; Long et al., 2000; van Dijk

unpublished data; present studyRana (Hylarana) taipehensis van Denburgh, 1909 Present studyRana (Pelophylax) lateralis Boulenger, 1887 Bourret, 1942; Long et al., 2000; present studyRana (Sylvirana) faber, new species Present studyRana (Sylvirana) mortenseni Boulenger, 1903 Present studyRhacophorus bipunctatus Ahl, 1927 Present studyRhacophorus bisacculus Taylor, 1962 van Dijk unpublished data; present studyTheloderma asperum (Boulenger, 1893) Present study

MICROHYLIDAE Gunther, 1858

Glyphoglossus molossus Gunther, 1869 Bourret, 1942; van Dijk unpublished dataKalophrynus interlineatus Blyth, 1855 Bourret, 1942; van Dijk unpublished data; present studyKaloula pulchra Gray, 1831 Flower, 1896; Long et al., 2000; van Dijk unpublished data;

Present studyMicryletta inomata (Boulenger, 1890) van Dijk unpublished data; present studyMicrohyla annamensis Smith, 1923 Present studyMicrohyla berdmorei (Blyth, 1856) Flower, 1896; Long et al., 2000; van Dijk; unpublished data;

Present studyMicrohyla butleri Boulenger, 1900 Present studyMicrohyla heymonsi Vogt, 1911 Bourret, 1942; Long et al., 2000; van Dijk unpublished data; I

Present studyMicrohyla ornata (Dumeril & Bibron, 1841) Boulenger, 1882 Flower, 1899; Long et al., 2000; van Dijk

unpublished data; present studyMicrohyla pulchra (Hallowell, 1861) Boulenger, 1882; Flower, 1899; Long et al., 2000; present study

i

479 J


were conducted by other researchers during the time of the Bourret, R., 1941. Notes herpetologiques sur l'Indochine fran~aise.Cardamom and Northeast Mondulkiri surveys reported here XXII. Reptiles et Batraciens re~us au Laboratoire des Sciences(WCS unpublished data). Unfortunately the species collected Naturelles de 1'Universite au cours de l' annee 1941. Descriptionduring these surveys have yet to be identified and therefore d'une espece et d'une variete nouvelles. Annexe du Bulletinthese data cannot be included in the national checklist de l'Instruction publique, Hanoi, 1941: 5-29.presented here. Undoubtedly the known amphibian fauna Bourret, R, 1942. Les Batraciens de l'Indochine. Instituteof Cambodia will expand rapidly over the next decade or so Oceanographique de 1'Indochine, Hanoi. x + 547 pp., 4 pl.as more areas of the country are explored for the first time .b b. I . I t It . h d th t th It f Daltry, J. C. & Chheang Dany, 2000. Reptiles. In: Daltry, J. C. &

Y 10 oglca survey eams. IS ope a e resu so, " ,th .. tI' I h ' b . fC b d . rt d h ' II F. Momberg (eds.), 2000. Cardamom Mountams Biodiversity

e ml a amp 1 Ian surveys 0 am 0 la repo e ere WI , ..d I. bl f d t. f f t h b . ld Survey 2000. Fauna & Flora International, Cambndge, UK.provl e a re la e oun a Ion or u ure researc to Ul 254 .upon. pp

Daltry, J. C. & F. Momberg (eds.), 2000. Cardamom MountainsBiodiversity Survey 2000. Fauna & Flora International,

ACKNOWLEDGEMENTS Cambridge, UK. 254 pp.

Deuve, Th" 2002. Nouveaux Carabus L. et Cychrus F, de ChineThe surveys for amphibians in both the Cardamom occidentale (Coleoptera, Carabidae). Coleopteres, 8: 239-246.Mountains and Northeast Mondulkiri. Pro;ince w~re Dring, J. C, M., 1987. Bornean tree frogs of the genus Philautusconducted as part of Fauna & Flora International s Indochma (Rhacophoridae). Amphibia-Reptilia, 8: 19-47.Programme in collaboration with the Department of Forestry, "and Wildlife (Ministry of Agriculture, Forestry and Dubol~, ~. & A, Ohler, 1998. A ne-:v species ,of Lept~brachlumFisheries), and the Department of Nature Conservation and (Vlbrlssaphora) from northern Vietnam, -:vrth a review ~f theProtection (Ministry of the Environment). We are indebted taxonom~ of the gen,u.s Leptobrachlum (Pelobatldae,to the Government of Cambodia for providing the Megophrymae). Dumerula, 4 (1): 1-32.

opportunity to work in Cambodia. We are grateful to all Dubois,A. &A. Ohler, 1999. Asian and Oriental toads of the Bufoour guides who were essential to our fieldwork, and melanostictus, Bufo scaber and Bufo stejnegeri groupsespecially to Kry Masphal for his assistance in the field in (Amphibia, Anura): a list of available and valid names and2000, and Jeremy Holden who collected many of the redescription of some name-bearing types. Journal of Southspecimens from the Cardamom Mountains, including those Asian Natural History, 4 (2): 133-180,

of the new Philautus species. Peter Paul van Dijk provided Dubois, A. & A. Ohler, 2000. Systematics of Fejervaryaconstructive advice and kindly granted access to unpublished limnocharis (Gravenhorst, 1829) (Amphibia, Anura, Ranidae)data. Alain Dubois and Miguel Vences gave useful comments and related species. 1. Nomenclatllral statlls and type-specimensto the manuscript. This is publication N°65 of the of the nominal species Rana limnocharis Gravenhorst, 1829."Programme Pluriformation Asie du Sud-Est. Publication Alytes, 18 (1-2): 15-50.N°64 see Deuve, 2002). Fei, L., C. Ye & Y. Huang, 1991. Key to Chinese Amphibia.

Chongqing, Editions of Sciences and Techniques. [iv] + 2 +364 pp, [In Chinese.]

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306 00 oglca oclety OJ non" .

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s we,., .atlona 10 Iverslty rospectus: d ' th M 1 P , 1 d S.f N b 1896ma e mea ay emnsu a an lam, rom ovem er

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Fontanel, J., 1972. Carton des Bioclimats. In Notice de la CarteBossuyt, F. & A. Dubois, 2001. A review of the frog genus du Cambodge. (Carte Internationale du Tapis Vegetal et de

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Boulenger, G. A., 1882. Catalogue of the Batrachia Gradentia s, Pondichery, Hors ser. 11:1-238.Caudata in the collection of the British Museum. London, Frost, D. R., 2000. Amphibian species of the world: an onlineTaylor & Francis. xvi + 503pp., 30 pl. reference, Version 2.20, http://research,arnnh.org/herpetology/

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Boulenger, G. A., 1908. A revision of the Oriental pelobatid Inger, R. F., 1970. A new species of frog of the genus Rana frombatrachians (Genus Megalophrys). Proceedings of the Thailand. Fieldiana: Zoology Memoires, 51: 169-174.Zoological Society of London, 1908: 407-430. Inger, R. F., 1999. Distribution of amphibians of southern Asia

Boulenger, G. A., 1920. A monograph of the South Asian, Papuan, and adjacent islands. In: Duellman, W. E. (ed.), Patterns ofMelanesian, and Australian frogs of the genus Rana. Records distribution of amphibians: A global perspective. John Hopkinsof the Indian Museum, 20: 1-126. University Press, Baltimore: 456-482.

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Le, T. T., W. J. Richardson, V. C. Le, H. M. Tran, Q. N. Tran, V. (Sumatra, Java, Borneo) with the definition of two species.S. Nguyen, A. L. Monastyrskii & J. C. Eames, 1999. A Alytes, 19 (1): 5-28.Feasability Study for the Establishment of Phong Dien (Thua, ...,Thien Hue Province) and Dakrong (Quang Tri Province) Welle~, H., 1998. The distribution of tiger,. leopard, elephant a~dNature Reserves, Vietnam. BirdLife International Vietnam wild cattle (gaur, banteng, buffalo, khtmg vor and kouprey) mProgramme, Hanoi. Cambodia. Cat Action Treasury/Wildlife Protection Office,

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..Material studied. -Megophrys (Xenophrys) major: BMNHMocquard~ R., 1904: Batrac.ie~s recu~llls par~. A. Pavle en 1868.7.3.98,1 adult male, Pegu, Myanmar; BMNH 1947.2.24.92-

Indochme. InPavle, A.,MlsslonPavle/ndo-Chme. /879-/895. 96 t fX h . J 1870 4 d It 1 1E d d' III R h h l 'h'. 11 d ' syn ypes 0 enop rys gigas erson, ,a u ma es,tu es Iverses. .ec erc es sur Istoue nature e e d 1 " 1 D . 1. I d. BMNH 18724 17431 1.. 11'1 d Ch.. 1 P . E L . 473 474 aut lema e, arJee lng, n la; ...,Juvem en 0- me onenta e. ans, rnest eroux. -" 1 D . I ' I d. MSNG 29066 1 d 1 1 S.kkilema e, arJee lng, n la; , aut ma e, I m,Momberg,F. &J. C. Daltry, 2000. General Introduction. In: Daltry, India; BMNH 1908.4.8.4-6,2 adult males, 1 juvenile female,

J. C. & F. Momberg (eds.), Cardamom. Mountains Bfodiversity Cherrapunji, Meghalaya, India; MNHN 1893.0514-0516, 1 adultSurvey 2000. Fauna & Flora International, Cambndge, UK. male, 1 sub-adult male, 1 juvenile female, Karin Bia-Po, Myanmar;

Ohler, A., 1997. Amphibiens. Unpublished draft, Paris, MNHN. 2 BMNH 1940.6.2.4-10, 1974.810,2 adult males, 6 adult females,pp. Pangnamdim, Triangle, Myanmar; MNHN 1987.2183, 1 juvenile

Ohler, A. & A. Dubois, 1999. The identity of Elachyglossa female, Doi Pui, Thailand; MNHN 1986.2184-2185,1 adult male,gyldenstolpei Anderson, 1916 (Amphibia, Ranidae), with 1 adult female, Siriphum, Doi Inthanon, Thailand; MNHNcomments on some aspects of statistical support to taxonomy. 1986.2186, 1 sub-adult male, Huai Fha Mon, Doi Inthanon,Zoologica Scripta, 28, (3-4): 269-279. Thailand; MNHN 0000,8182, lectotype of Megophrys longipes

maosonensis Bourret, 1937, 1 juvenile female, Maoson, Vietnam;Ohle~, A.,.O. MarquIs, S. S,,:,an & S. GrosJean, 2000. ~mph~blan MNHN 1938.0096-0097 0000.8181 aralectot esofMe 0 h sdiversity of the Hoang Lien Nature Reserve (Lao Cal Provmce, ., ' , p yp g V rynorthern Vietnam), with a description of two new species. lo~glpes maosonensls Bourret, 1937, 3 adult males, Maoson,Herpetozoa, 13 (1/2): 71-87. Vietnam; MNHN 1938.0098-0099, paralectotypes of Megophrys.longipes maosonensis Bourret, 1937, 1 adult male, 1 sub-adult

Robichaud, W. & B. L. Stuart, 1999. Summary of Saola, male SaPa Vietnam' MNHN 0000.9179-8180 paralectotypes ofHerpetological and Wildlife Trade Studies in Nakai-Nam Theun M ' h '

I .' . Bo t 193'7 1 ad It " male' ., egop rys onglpes maosonensls urre , , u Ie ,

NBCA and the Proposed Nam Theun Extension. Wildlife 1 .. 1 " 1 S P V. t RMNH 4770 1 d It 1C . S .L P v,. Juvem e lema e, a a, Ie nam; , a u ma e,onservatlon oclety aos rogram, lentlane. A V. MSNG29453 1 d 1 " 1 1..

1 alnnam, letnam; , aut lema e, Juvem erne,Smith, M. A., 1922a. The frogs allied to Rana doriae. Journal of 1 juvenile female, Mount Karin, Myanmar, colI. L. Fea, 1888.

the Natural History Society of Siam, 4: 215-225. Megophrys (Xenophrys) robusta: BMNH 1947.2.25.19, syntype,

Smith, M. A., 1922b. On a collection of reptiles and batrachians Darjeeling, India; BMNH 1872.4.17.430, Darjeeling, India.from Siam and Indo-China (No.1). Journal of the Natural Limnonectes gyldenstolpei: BMNH 1947.2.1.97-99, 1947.2.2.1,History Society of Siam, 4: 203-214, pl. 8. 1947.2.2.3-5, syntypes, Khao Sebab, Chantabun, Thailand; BMNH

Smith, M. A., 1930. The Reptilia and amphibia of the Malay 1947.2.2.10, syntype, Meh Song, fore~t near Pr~e, Thailand;Peninsula. Bulletin of the Raffles Museum, 3: i-xviii + 1-149. BMNH 1947.2.2.11, syntype, HupBon, Snracha, Thailand; MNHN

1997.4149-4150, Bokeo, Laos.Steinheime~, F. D., J. C. Eames, M. Chandamony & R. Bansok, Paafasciculispina: MNHN 1989.0706-0710, topotypes, Kao Soi2000. Buds. In: Daltry, J. C. & F. Momberg (eds.), Cardamoms D Ch t Ch t b . Th 'I dB . d .. S 2000 87 9 8 F & Fl ao, angwa an a un, aI an .10 Iverslty urvey , pp. -.auna ora R '. BMNH 19741957-19741968 9 d 1 1 3I t t. 1 C b .d 254 ana mortensen!, .., aut ma es,n erna lona, am n ge. pp. d 1 al . Th ' I B 194 2 51aut fern es, Ban Pipeng, al and; MNH 7.2.., syntype,

Stuart, B. L., 1999. Amphibians and Reptiles. In: Duckworth, J. adult male, Koh Chang, Thailand; BMNH 1915.8.14.14,

W., R. E. Salter & K. Khounboline (compilers), Wildlife in 1919.3.28.1-2, 1921.2.12.1, 1 adult male, 3 adult females,Lao PDR: ~99~ Status Report. Pp. 43-67. IUCN, WCS, topotypes, Koh Chang, Thailand.CPA WM, Vientiane. Rana nigrovittata: BMNH 1947.2.2.93, lectotype, ZSI 2685,

Taylor, E. H., 1962. The amphibian fauna of Thailand. University paralectotype, 2 adult males, ZSI 2773, paralectotype, adult female,of Kansas Science Bulletin, 43(8), 265-599. Mergui, Myanmar.

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